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NOTES ON THE LIFE CYCLE AND NATURAL HISTORY OF
BUTTERFLIES OF EL SALVADOR. VIII. ARCHAEOPREPONA
ANTIMACHE GULINA, SIDERONE MARTHESIA, ZARETIS
CALLIDRYAS AND CONSUL ELECTRA (NYMPHALIDAE)
Alberto Muyshondt 101 Avenida Norte #322, San Salvador, El Salvador
This article completes a series dealing with what my sons and I have learned about the early stages, larval food plants, and adult behavior of the Charaxinae found in El Salvador. As far as we know, the remaining local species of this group for which we have been unable to find the early stages, even though we have collected adults, are Anaea (Anaea) aidea Guerin-Meneville, A. (Memphis) nobilis Bates and A. (M.) glycerium Doubleday. Another species of Archaeoprepona occurs here, but we unfortunately have been unable to capture an adult to have it determined. All these species are very scarce locally; only one or two specimens of some are known in local collections (including ours).
Life Cycles
Archaeoprepona antimache gulina Friihstorfer
This species is like A. demophon centralis Friihstorfer (Muyshondt, 1976) except for the following points:
Larva (Figs. 2-6). Dark brown dorsally during 1st and 2nd instars; 3rd instar lighter brown with a whitish spiracular area on 7th and 9th abdominal segments; 5th instar larva whitish dorsally from 3rd-9th abdominal segments, rest of body brown. During all instars the mesal thoracic prominence is larger than in respective instars of A. d. centralis.
Prepupa (Fig. 7). Prepupa changes to yellow instead of green.
Pupa (Figs. 8,9). Pupa slightly more slender laterally than pupa of A. d. centralis.
Adult (Figs. 10, 11). Same shape as A. d. centralis but colored triangle on dorsal surface of forewing and band on dorsum of hindwing are definitely iridescent light blue instead of iridescent greenish-blue. Ventral surfaces of wings mostly light gray instead of light brown with a complicated array of darker zones and black lines, more abundant than in A. d. centralis. The wingspan and developmental time are about the same as A. d. centralis.
Siderone marthesia Cramer
In all respects, this species is like Zaretis itys Cramer (Muyshondt, 1973b) but for the following:
Egg. Considerably larger, but same shape and color.
Larva (Figs. 12-16). Corresponding instars about 1% times larger, lighter brown than Z. itys; caudal end rounded instead of bifid. Later instars with a white patch formed by short and thin spinulets subdorsally on mesothorax. Head horns very thick and knobby; head and its horns covered with short setae.
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Figs. 1-6. Archaeoprepona antimache gulina. 1, Egg, width about 2.5 mm; 2, second instar larva, about 13 mm long; 3, third instar larva, about 30 mm long; 4, fourth instar, about 47 mm long; 5, head of fifth instar larva, note lateral projections; 6, fifth instar larva, about 80 mm long.
Pupa (Figs. 18-20). Bigger than but about the same shape as pupa of Z. ittjs. Green with dark markings along the spiracular zone and on the lateral edges of the wingcases. The latter have a marked indentation, lacking in Z. itys.
Adults (Figs. 21, 22). Sexually dimorphic. Wing of male: costal margin convex, apex pointed, outer margin S-shapcd, tornus rounded and inner margin of forcwing straight. Costal margin of hindwing strongly rounded basally, outer angle and margin rounded, anal angle slightly projecting, inner margin S-shaped. Female: larger with costal margin of forewing less convex than in the male, apex without a marked point, outer margin slightly convex, forewing not S-shaped. Hindwing as in the male. Dorsal color pattern of the male: forewing dark brown, almost black, with a slanting red bar from the mid-costal margin almost to the tornus and a red triangle basally, leaving a black band in between. Male hindwing mostly black or very dark brown with red markings of variable size starting from mid-costal margin, markings sometimes reduced to two or three spots or forming a band running toward the anal angle. Apex of front wing in some individuals red-tinged and with a very small white spot near costal margin of the subapical area. Females have same color pattern, but lighter brown with two subapical white spots on forewing and red markings of hindwing greatly reduced. Ventral wing surfaces of both sexes very
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Figs. 7-11. Archaeoprepona antimache gulina. 7, Prepupa, note tails located on either side of twig; 8, pupa, dorsal view, about 42 mm long; 9, pupa, ventral view; 10, male, dorsal view, wingspan about 80 mm; 11, male, ventral view.
similar but darker in male: ground color dark brown with darker lines resembling leaf veins and lighter zones, the latter more evident on hindwing. Apex of forewing with some white sprinkling. The whole of ventral wing surfaces gives silvery sheen at certain angles. Thorax reddish above, yellowish below; abdomen brown. Antennae light brown; palpi and proboscis yellow. Wingspan of female about 7.0 cm, male 5.5 cm.
Zaretis callidryas R. Felder
A second instar of Z. callidryas was collected and at first thought to be a different morph of the common Z. itys. The larva was reared on Casearia silvestris and C. nitida. During the later instars differences were noticed and photographed:
Larva (Fig. 23). Abdomen lighter dorsally than Z. itys; head smoother, without the typical Z. itys spines and the horns incurved posteriorly.
Pupa. Closely resembles the green morph of Z. itys.
Adult. The obtained adult was a tailed female. Its shape similar to Z. itys except for tail on M-3 of hindwing and less acute anal angle. Dorsal color predominantly
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Figs. 12-14. Siderone marthesia. 12, Fourth instar larva, about 32 mm long; 13, same, dorsal view; 14, fifth instar larva, two days after molting, about 44 mm long.
Fig. 15. Left, Zaretis itys, fifth instar larva; right, Siderone marthesia, fifth instar larva, about 62 mm long. Compare caudal end of Zaretis with Siderone.
Figs. 16-17. Close-up of larval heads: 16, Siderone and 17, Zaretis.
white with pearly sheen; apex of forewing faintly gray. Outer margin of hindwing with submarginal row of small faint brown spots. Ventially wings yellowish-white with faint markings as in Z. itys. Wingspan about 5 cm.
Consul electra Westwood
In its early stages, behavior and selection of food plant, this species is similar to C. fabius (Cramer) (Muyshondt, 1974) except for the following noticeable differences :
Larva (Figs. 26, 27). During 5th instar, head predominantly green with black near the horns and yellow frontal bands. In C. fabius the head is mostly black with yellow frontal bands,
Pupa (Figs. 28-30). Green and brown pupal morphs occur. Slightly wider laterally than in C. fabius; shows two dark spots dorsally on thorax.
Adult (Figs. 31-38). Adults not showing marked sexual dimorphism, but are amazingly polymorphic. Apex of forewing more or less elongate and acuminate; M-3
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Figs. 18-22. Siderone marthesia. 18, Pupa, dorsal view, about 25 mm long; 19, pupa, lateral view; 20, pupa, ventral view; 21, male, dorsal view, wingspan about 55 mm; 22, male, ventral view.
of hindwing more or less elongate and spatulate, anal angle sometimes forming second tail. Dorsal and ventral colors more or less dark with some variations in color pattern. Variations in shape and color exist between individuals reared simultaneously, thus, not attributable to seasonal variation. Wingspan of female about 7.0 cm, male 5.5 cm.
Natural History
Archaeoprepona a. gulina behaves during its early stages and as an adult very much like Prepona omphale octavia (Muyshondt, 1973a) and A. demophon centralis (Muyshondt, 1976). It looks very much like the latter except for variations in color. The larva of A. a. gulina also reacts in a menacing manner by lifting the forked tail which is opened and closed scissor-like when prodded. During the later instars
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Figs. 23-25. Zaretis callidryas. 23, Fifth instar larva, about 40 mm long, note incurved horns, compare with Z. itys, Fig. 15; 24, female (tailed morph), dorsal view, wingspan about 50 mm; 25, female, ventral view.
it also has a reptilian aspect when viewed dorsally. This is due to a prominent dorsal projection which covers the head. The lateral aspects of the dorsal projection bear false eyes which enhance the effect. This prominence also is retractable.
A. a. gulina larvae feed on a Laurinaceae tree determined by Lie. Flores, Universidad Nacional, as Nectandra sinuata, which is the same food plant used by the larvae of Anaea morvus hoisduvali Cornstock (Muyshondt, 1975). Actually it was while searching for A. m. hoisduvali eggs that the first eggs of A. a. gulina were found. The eggs of the latter species are also heavily parasitized by chalcidid wasps, and the small larvae are preyed upon by spiders. A. a. gulina occupies the same habitat as A. demophon centralis.
Siderone marthesia and Zaretis callidryas behave like Z. itys (Muyshondt, 1973b) and share its food plants: Casearia sylvestris and C. nitida although Zaretis itys appears to prefer the latter and Siderone marthesia and Zaretis callidryas the former. Both S. marihesia and Z. callidryas have been collected at an altitude of 1,000 m. Z. itys has a wider range. The females of S. marthesia are very seldom seen. Most of our observations have been of adult males; they have definite hill-topping behavior between 1030-1600 hr. They usually perch on the top branches of trees, and it is quite a feat to net them. Luckily they are attracted by fermenting banana baits, but we have never found females at the baits. However, Descimon et al. (1973) report several captures of females at baits near Colima, Mexico. We are unable to
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Figs. 26-34. Consul electra. 26, Fifth instar larva, about 37 mm long; 27, close-up of larval head; 28, pupa, dorsal view, about 20 mm long; 29, pupa, lateral view; 30, pupa, ventral view. Emergence of adult: 31-34, sequenced from left to right.
comment on the behavior of adult Zaretis callidryas because the only specimen we have seen is the one reared in our insectary.
The behavior of the early stages and adult of Consul electra resemble those of Anaea (Consul) fabius Cramer (Muyshondt, 1974). The larvae of both use a number of species of Piperaceae as food plants. C. electra has been collected at higher altitudes than fabius (500-2,600 m). Both
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Figs. 35-38. Consul electra. 35, Female, dorsal view, wingspan about 70 mm; 36, female, ventral view; 37, male (acute apex morph), dorsal view, wingspan about 55 mm; 38, male, ventral view.
species can be found simultaneously feeding on the same plants where their ranges merge at an altitude of 500-1,200 m. The adults do not visit flowers, but feed on fermenting fruits and excrement on the ground and on sap from tree cavities. The males are pugnacious and perch on shrubs at a height of 2-4 m. With C. fabius, C. electra is one of the slowest flyers among the local Charaxinae. The larvae and pupae are often parasitized by tachinid flies and chalcidid wasps.
Discussion
Miiller (1886) described the life cycles of various Archaeoprepona (under the generic name of Prepona), including A. amphimachus Fabricius on Laurinaceae (probably Nectandra spp.), A. catachlora Staudinger on Mollinedia brasiliensis Jul. (Monimaceae) and A. demo-phon extincta Staudinger on Dugetia lanceolata St. Hilaire (Anonaceae).
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His descriptions conform closely with our own observations of A. demo-phon centralis (Muyshondt, 1976) and A. antimache gulina except for the eggs which he describes as having 'Very fine and crowded longitudinal ribs" which we have been unable to detect, even under 15 X magnification. Miiller also described the early stages of Siderone ide Hiibner and S. strigosus Gmelin and claimed both species fed on Cas-earia sylvestris. However, Siderone strigosus, according to Comstock (1961), is a synonym of Anaea (Zaretis) itys. Furthermore, because Miiller states that the early stages of "Siderone strigosus' are "as ide" it appears that his descriptions actually deal with two species of Zaretis.
After studying species of both genera (Zaretis itys (Muyshondt, 1973b), Z. callidryas, and Siderone marthesia), we are certain that there are notable differences between the early stages of the larvae of the two genera that would not escape the attention of an investigator. The only part in Miiller's description of "ide" that would fit a Siderone pupa is "wing edge and stigma brown," yet he does not mention a drastic indentation (as seen dorsally and ventrally) at the brown mark on the wingcase of the pupa he described.
According to Comstock (1961), the only information about the early stages of A. (Siderone) marthesia is Miiller's, however, we are certain that Miiller's description was not of a species of Siderone. In Rydon (1971) there are some good painted illustrations of the larvae of S. marthesia and S. nemesis and their respective pupae copied from the original drawings of Miss M. E. Fontaine which were published by the Royal Entomological Society around 1910. However, Miss Fontaine apparently did not write any descriptions.
We are unaware of any previous publications on the early stages of Zaretis callidryas. The adult is figured in Seitz (1924) as an Anaea and is not likened to its close relative, Zaretis itys. Even in Lewis (1973) the tailless form of Z. callidryas is presented as an Anaea. The same is true with Consul electra. Seitz (1924) and Lewis (1973) present it as Anaea electra and do not relate it to its close relative, Consul fabius, which they picture as Protogonius hippona Fabricius.
We hope our descriptions and illustrations of the species we have studied will help the experts clarify and settle the apparent congeneric questions of the Charaxinae.
Acknowledgments
We are greatly thankful to Drs. A. H. B. Rydon and H. Descimon for supplying us with a number of publications that helped document the present article; to Drs. James G. Sternburg and G. L. Godfrey for dedi-
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eating part of their valuable time to reading, criticizing, and suggesting improvements for the manuscript; to Viktor Hellebuyck for sharing his own observations of the species mentioned; and to Alberto, Jr., Marilynn and Pierre, members of the family, for their enthusiastic help.
Literature Cited
Comstock, W. P. 1961. Butterflies of the American Tropics, the genus Anaea,
(Lepidoptera, Nymphalidae). The American Museum of Natural History, N.Y.
214 p. Descimon, H., J. M. de Maeght & J.-R. Stoffel. 1973. Contribution a I'etude
des Nymphalides neotropicales. Description de trois nouveaux Prepona mexi-
cains. Alexanor 8: 155-159. Hemming, F. 1967. The generic names of the butterflies and their type-species.
(Lepidoptera: Rhopalocera). Bull. British Mus. (Nat. Hist.), Entomology
Suppl. 9. 509 p. Lewis, H. L. 1973. Butterflies of the World. Harrap Books, London; and Follett,
Chicago, xvi + 312 p. Muller, W. 1886. Sudamerikanische Nymphalidenraupen. Versuch eines natiir-
lichen Systems der Nymphaliden. Zool. Jahrb. Zeitschr. Syst., Geogr., Biol, der
Thiere 1: 417-678. Muyshondt, A. 1973a. Notes on the life cycle and natural history of butterflies
of El Salvador. I. Prepona omphale octavia (Nymphalidae). J. Lepid. Soc. 27:
210-219. ----------. 1973b. Notes on the life cycle and natural history of butterflies of El
Salvador. II. Anaea (Zaretis) itys (Nymphalidae). J. Lepid. Soc. 27: 294-302. ----------. 1974. Notes on the life cycle and natural history of butterflies of El
Salvador. III. Anaea (Consul) fabius (Nymphalidae). J. Lepid. Soc. 28:
81-89. ----------. 1975. Notes on the life cycle and natural history of butterflies of El
Salvador. V. Anaea (Memphis) morvus boisduvali (Nymphalidae). J. Lepid.
Soc. 29: 32-39. ----------. 1976. Notes on the life cycle and natural history of butterflies of El
Salvador. VII. Archaeoprepona demophon centralis (Nymphalidae). J. Lepid.
Soc. 30: 23-32. Rydon, A. H. B. 1971. The systematics of the Charaxidae (Lepidoptera: Nymph-
aloidea). Ent. Rec. J. Var. 83: 219-233, 283-287, 310-316, 336-341, 384-388. Seitz, A. 1924. Macrolepidoptera of the World. Vol. 5. Stuttgart.